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A pupa (from Latin pupa 'doll'; pl.: pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages thereof being egg, larva, pupa, and imago. The processes of entering and completing the pupal stage are controlled by the insect's hormones, especially juvenile hormone, prothoracicotropic hormone, and ecdysone. The act of becoming a pupa is called pupation, and the act of emerging from the pupal case is called eclosion or emergence.

The pupae of different groups of insects have different names such as chrysalis for the pupae of butterflies and tumbler for those of the mosquito family. Pupae may further be enclosed in other structures such as cocoons, nests, or shells.^[1]

Position in life cycle

The pupal stage follows the larval stage, or in some cases a prepupal stage, and precedes adulthood (imago) in insects with complete metamorphosis. The pupa is a non-feeding, usually sessile stage, or highly active as in mosquitoes. It is during the pupal stage that the adult structures of the insect are formed while the larval structures are broken down. The adult structures grow from imaginal discs.^[2]

Duration

The pupal stage may last weeks, months, or even years, depending on temperature and the species of insect.^[3]^[4] For example, the pupal stage lasts eight to fifteen days in monarch butterflies.^[5] The pupa may enter dormancy or diapause until the appropriate season to emerge as an adult insect. In temperate climates pupae usually stay dormant during winter, while in the tropics pupae usually do so during the dry season.

Eclosion of Papilio dardanus

Emergence

Insects emerge (eclose) from pupae by splitting the pupal case. Most butterflies emerge in the morning. In mosquitoes, the emergence is in the evening or night. In fleas, the process is triggered by vibrations that indicate the possible presence of a suitable host. Prior to emergence, the adult inside the pupal exoskeleton is termed pharate. Once the pharate adult has eclosed from the pupa, the empty pupal exoskeleton is called an exuvia; in most hymenopterans (ants, bees and wasps) the exuvia is so thin and membranous that it becomes "crumpled" as it is shed. Measuring the timing of this emergence is of interest to chronobiologists because the process is regulated by circadian clocks in many species, necessitating different assays to measure eclosion timing.

Pupal mating

In a few taxa of the Lepidoptera, especially Heliconius, pupal mating is an extreme form of reproductive strategy in which the adult male mates with a female pupa about to emerge, or with the newly moulted female; this is accompanied by other actions such as capping of the reproductive system of the female with the sphragis, denying access to other males, or by exuding an anti-aphrodisiac pheromone.^[6]^[7]

Defense

Pupae are usually immobile and are largely defenseless. To overcome this, pupae often are covered with a cocoon, conceal themselves in the environment, or form underground.^[8] Some species of Lycaenid butterflies are protected in their pupal stage by ants. Another means of defense by pupae of other species is the capability of making sounds or vibrations to scare potential predators. A few species use chemical defenses including toxic secretions. The pupae of social hymenopterans are protected by adult members of the hive.

Types

Based on the presence or absence of articulated mandibles that are employed in emerging from a cocoon or pupal case, the pupae can be classified in to two types:^[9]

Decticous pupa – a pupa with articulated mandibles. Examples are pupae of the orders Neuroptera, Mecoptera, Trichoptera and few Lepidoptera families.
Adecticous pupa – a pupa without articulated mandibles. Examples include the orders Strepsiptera, Coleoptera, Hymenoptera, Diptera and Siphonaptera.

Based on whether the pupal appendages are free or attached to the body, the pupae can be classified as one of three types:

Exarate pupa – appendages are free and are not usually encapsulated within a cocoon. Decticous pupae are always exarate; some adecticous pupae are as well. (Neuroptera, Trichoptera, Cyclorrhapha of Dipterans, Siphonaptera, most Coleoptera, Hymenoptera, and few Lepidoptera).
Obtect pupa – appendages are attached closely to the body and are commonly encapsulated within a cocoon. Some adecticous pupa are obtect forms. (Most Lepidoptera, Nematocera and Brachycera of Dipterans, Staphylinidae and Chrysomelidae Coleopterans, many Chalcidoidea Hymenopterans)
Coarctate pupa – enclosed in a hardened cuticle of the penultimate larval instar called a puparium. However, the pupa itself is of the exarate adecticous pupal form. (Cyclorrhapha of Dipterans).

Chrysalis

A chrysalis (Latin: chrysallis, from Ancient Greek: χρυσαλλίς, chrysallís, plural: chrysalides, also known as an aurelia) or nympha is the pupal stage of butterflies. The term is derived from the metallic–gold coloration found in the pupae of many butterflies, referred to by the Ancient Greek term χρυσός (chrysós) for gold.

When the caterpillar is fully grown, it makes a button of silk which it uses to fasten its body to a leaf or a twig. Then the caterpillar's skin comes off for the final time. Under this old skin is a hard skin called a chrysalis.^[10]

Because chrysalises are often showy and are formed in the open, they are the most familiar examples of pupae. Most chrysalides are attached to a surface by a Velcro-like arrangement of a silken pad spun by the caterpillar, usually cemented to the underside of a perch, and the cremastral hook or hooks protruding from the rear of the chrysalis or cremaster at the tip of the pupal abdomen by which the caterpillar fixes itself to the pad of silk. (Gr. kremastos 'suspended')^[11]

Like other types of pupae, the chrysalis stage in most butterflies is one in which there is little movement. However, some butterfly pupae are capable of moving the abdominal segments to produce sounds or to scare away potential predators. Within the chrysalis, growth and differentiation occur.^[12] The adult butterfly emerges (ecloses) from this and expands its wings by pumping haemolymph into the wing veins.^[13] Although this sudden and rapid change from pupa to imago is often called metamorphosis, metamorphosis is really the whole series of changes that an insect undergoes from egg to adult.

When emerging, the butterfly uses a liquid, sometimes called cocoonase, which softens the shell of the chrysalis. Additionally, it uses two sharp claws located on the thick joints at the base of the forewings to help make its way out.^[14] Having emerged from the chrysalis, the butterfly will usually sit on the empty shell in order to expand and harden its wings. However, if the chrysalis was near the ground (such as if it fell off from its silk pad), the butterfly would find another vertical surface to rest upon and harden its wings (such as a wall or fence).

Moth pupae are usually dark in color and either formed in underground cells, loose in the soil, or their pupa is contained in a protective silk case called a cocoon. The pupa of some species such as the hornet moth develops sharp ridges around the outside called adminicula that allow the pupa to move from its place of concealment inside a tree trunk when it is time for the adult to emerge.^[15]

Pupa, chrysalis, and cocoon are frequently confused, but are quite distinct from each other. The pupa is the stage between the larva and adult stages. The chrysalis generally refers to a butterfly pupa although the term may be misleading as there are some moths whose pupae resembles a chrysalis, e.g.: the plume winged moths of the family Pterophoridae and some geometrid moths. A cocoon is a silk case that the larvae of moths, and sometimes other insects, spin around the pupa.

Cocoon

A cocoon is a casing spun of silk by many moths and caterpillars,^[16] and numerous other holometabolous insect larvae as a protective covering for the pupa.

Cocoons may be tough or soft, opaque or translucent, solid or meshlike, of various colors, or composed of multiple layers, depending on the type of insect larva producing it. Many moth caterpillars shed the larval hairs (setae) and incorporate them into the cocoon; if these are urticating hairs then the cocoon is also irritating to the touch. Some larvae attach small twigs, fecal pellets or pieces of vegetation to the outside of their cocoon in an attempt to disguise it from predators. Others spin their cocoon in a concealed location—on the underside of a leaf, in a crevice, down near the base of a tree trunk, suspended from a twig or concealed in the leaf litter.^[17] Contrary to popular belief, larvae do not completely liquify inside the cocoon.^[18]

The silk in the cocoon of the silk moth can be unraveled to harvest silk fibre which makes this moth the most economically important of all lepidopterans. The silk moth is the only completely domesticated lepidopteran; it does not exist in the wild.

Insects that pupate in a cocoon must escape from it, and they do this either by the pupa cutting its way out, or by secreting enzymes, sometimes called cocoonase, that soften the cocoon. Some cocoons are constructed with built-in lines of weakness along which they will tear easily from inside, or with exit holes that only allow a one-way passage out; such features facilitate the escape of the adult insect after it emerges from the pupal skin.

Puparium

Some pupae remain inside the exoskeleton of the final larval instar and this last larval "shell" is called a puparium (plural, puparia). Flies of the group Muscomorpha have puparia, as do members of the order Strepsiptera, and the Hemipteran family Aleyrodidae.^[19]

Gallery

Rhynchophorus ferrugineus pupa
Fruit fly (Drosophila melanogaster) pupa
An emperor gum moth caterpillar spinning its cocoon
Luna moth cocoon and pupa
Assortment of luna moth cocoons
Luna moth emerging from pupa within silk cocoon
Luna moth pupa removed from cocoon
Chrysalis of Gulf fritillary
Pupation of Aglais io
Monarch butterfly chrysalis
Specimen of an eclosing Dryas iulia butterfly
Pupae of Japanagromyza inferna, a gall fly, in gall of Centrosema virginianum
Pupa of cabbage looper
Pupa of baron butterfly Euthalia aconthea
Papilio polytes common Mormon pupa on curry leaves.
Dog-day cicada emerging

References

^ Borror, D. J.; DeLong, Dwight M.; Triplehorn, Charles A. (2004). Introduction to the Study of Insects (Sixth ed.). New York: Holt, Rinehart & Winston. ISBN 0-03-096835-6.
^ Aldaz, Silvia; Escudero, Luis M. (2010). "Imaginal discs". Current Biology. 20 (10): R429–R431. Bibcode:2010CBio...20.R429A. doi:10.1016/j.cub.2010.03.010. PMID 20504747.
^ Nielsen, Erik Tetens, and J. St Haeger. "Pupation and emergence in Aedes taeniorhynchus (Wied.)." Bulletin of Entomological Research 45.4 (1954): 757–768.
^ Elliott, J. M. "Temperature‐related fluctuations in the timing of emergence and pupation of Windermere alder‐flies over 30 years." Ecological Entomology 21.3 (1996): 241–247.
^ University of Minnesota Extension, Monarch Lab. "Monarch Life Cycle". Archived from the original on 7 July 2018. Retrieved 20 September 2017.
^ Preston–Mafham, Rod; Preston–Mafham, Ken (1993). The Encyclopedia of Land Invertebrate Behaviour (Illustrated ed.). MIT Press. p. 113. ISBN 978-0-262-16137-4. Retrieved 16 November 2010.
^ Boggs, Carol L.; Watt, Ward B.; Ehrlich, Paul R. (2003). Butterflies: Ecology and Evolution Taking Flight (Illustrated ed.). University of Chicago Press. p. 739. ISBN 978-0-226-06318-8. Retrieved 16 November 2010.
^ Chapman, R. F. (Reginald Frederick) (2013). The insects : structure and function. Cambridge University Press. p. 419. ISBN 978-0-521-11389-2. OCLC 794624696.
^ "Types of Pupa". Agri Info. Archived from the original on 4 February 2017. Retrieved 28 April 2016.
^ Darby, Gene (1958). What is a Butterfly. Chicago: Benefic Press. p. 19.
^ "Academic Dictionaries and Encyclopedias". Archived from the original on 2013-05-25. Retrieved 2013-01-20.
^ Lowe, Tristan; Garwood, Russell P.; Simonsen, Thomas; Bradley, Robert S.; Withers, Philip J. (2013). "Metamorphosis revealed: three dimensional imaging inside a living chrysalis". Journal of the Royal Society Interface. 10 (84). 20130304. doi:10.1098/rsif.2013.0304. PMC 3673169. PMID 23676900.
^ AMNH Archived December 7, 2006, at the Wayback Machine Accessed December 2006
^ The Entomologist
^ Newland, D.E. "Eclosion mechanics, mating and ovipositing behaviour of Sesia apiformis". Entomologist's Gazette.
^ Darby, Gene (1958). What is a Butterfly. Chicago: Benefic Press. p. 41.
^ Scoble, Malcolm J. (1992). The Lepidoptera: Form, Function and Diversity. Oxford: Oxford University Press. ISBN 0-19-854031-0.
^ Guarino, Ben (September 19, 2017). "Mutant butterflies reveal the genetic roots of colorful wings". The Washington Post. Archived from the original on September 19, 2017. Retrieved July 6, 2024.
^ Gullan, P. J.; Cranston, P. S. (2000). The Insects: An Outline of Entomology (2nd ed.). London: Blackwell. ISBN 0-632-05343-7.

External links

Pupa photos plus species descriptions at Insecta.pro
Silk worm life cycle photos (archived 8 November 2012)

[1] Borror, D. J.; DeLong, Dwight M.; Triplehorn, Charles A. (2004). Introduction to the Study of Insects (Sixth ed.). New York: Holt, Rinehart & Winston. ISBN 0-03-096835-6.

[AldazEscudero2010-2] Aldaz, Silvia; Escudero, Luis M. (2010). "Imaginal discs". Current Biology. 20 (10): R429–R431. Bibcode:2010CBio...20.R429A. doi:10.1016/j.cub.2010.03.010. PMID 20504747.

[3] Nielsen, Erik Tetens, and J. St Haeger. "Pupation and emergence in Aedes taeniorhynchus (Wied.)." Bulletin of Entomological Research 45.4 (1954): 757–768.

[4] Elliott, J. M. "Temperature‐related fluctuations in the timing of emergence and pupation of Windermere alder‐flies over 30 years." Ecological Entomology 21.3 (1996): 241–247.

[5] University of Minnesota Extension, Monarch Lab. "Monarch Life Cycle". Archived from the original on 7 July 2018. Retrieved 20 September 2017.

[Preston-Mafham_&_Preston-Mafham_(1993)-6] Preston–Mafham, Rod; Preston–Mafham, Ken (1993). The Encyclopedia of Land Invertebrate Behaviour (Illustrated ed.). MIT Press. p. 113. ISBN 978-0-262-16137-4. Retrieved 16 November 2010.

[Boggs,_Watt_&_Ehrlich_(2003)-7] Boggs, Carol L.; Watt, Ward B.; Ehrlich, Paul R. (2003). Butterflies: Ecology and Evolution Taking Flight (Illustrated ed.). University of Chicago Press. p. 739. ISBN 978-0-226-06318-8. Retrieved 16 November 2010.

[8] Chapman, R. F. (Reginald Frederick) (2013). The insects : structure and function. Cambridge University Press. p. 419. ISBN 978-0-521-11389-2. OCLC 794624696.

[9] "Types of Pupa". Agri Info. Archived from the original on 4 February 2017. Retrieved 28 April 2016.

[10] Darby, Gene (1958). What is a Butterfly. Chicago: Benefic Press. p. 19.

[11] "Academic Dictionaries and Encyclopedias". Archived from the original on 2013-05-25. Retrieved 2013-01-20.

[Lowe-12] Lowe, Tristan; Garwood, Russell P.; Simonsen, Thomas; Bradley, Robert S.; Withers, Philip J. (2013). "Metamorphosis revealed: three dimensional imaging inside a living chrysalis". Journal of the Royal Society Interface. 10 (84). 20130304. doi:10.1098/rsif.2013.0304. PMC 3673169. PMID 23676900.

[13] AMNH Archived December 7, 2006, at the Wayback Machine Accessed December 2006

[14] The Entomologist

[15] Newland, D.E. "Eclosion mechanics, mating and ovipositing behaviour of Sesia apiformis". Entomologist's Gazette.

[16] Darby, Gene (1958). What is a Butterfly. Chicago: Benefic Press. p. 41.

[17] Scoble, Malcolm J. (1992). The Lepidoptera: Form, Function and Diversity. Oxford: Oxford University Press. ISBN 0-19-854031-0.

[18] Guarino, Ben (September 19, 2017). "Mutant butterflies reveal the genetic roots of colorful wings". The Washington Post. Archived from the original on September 19, 2017. Retrieved July 6, 2024.

[19] Gullan, P. J.; Cranston, P. S. (2000). The Insects: An Outline of Entomology (2nd ed.). London: Blackwell. ISBN 0-632-05343-7.

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@@ Line 3: / Line 3: @@
 [[Image:Cetoine global.jpg|thumb|upright=1.6|Pupa of the rose chafer beetle, ''[[Cetonia aurata]]'']]
 [[Image:Mosquito Pupa.jpg|thumb|Tumbler (pupa) of a mosquito. Unlike most pupae, tumblers can swim around actively.]]
-A '''pupa''' ({{lang-la|pupa}}, "doll"; plural: ''pupae'') is the life stage of some [[insect]]s undergoing transformation between immature and mature stages. Insects that go through a pupal stage are [[Holometabolism|holometabolous]]: they go through four distinct stages in their life cycle, the stages thereof being egg, [[larva]], pupa, and [[imago]]. The processes of entering and completing the pupal stage are controlled by the insect's hormones, especially [[juvenile hormone]], [[prothoracicotropic hormone]], and [[ecdysone]]. The act of becoming a pupa is called pupation, and the act of emerging from the pupal case is called eclosion or emergence.
+A '''pupa''' ({{ety|la|pupa|doll}}; {{plural form}}: '''pupae''') is the life stage of some [[insect]]s undergoing transformation between immature and mature stages. Insects that go through a pupal stage are [[holometabolous]]: they go through four distinct stages in their life cycle, the stages thereof being egg, [[larva]], pupa, and [[imago]]. The processes of entering and completing the pupal stage are controlled by the insect's hormones, especially [[juvenile hormone]], [[prothoracicotropic hormone]], and [[ecdysone]]. The act of becoming a pupa is called pupation, and the act of emerging from the pupal case is called eclosion or emergence.
-The pupae of different groups of insects have different names such as [[#Chrysalis|''chrysalis'']] for the pupae of [[butterflies]] and ''tumbler'' for those of the [[mosquito]] family. Pupae may further be enclosed in other structures such as [[#Cocoon|cocoons]], [[nest]]s, or [[Animal shell|shells]].<ref>{{cite book |last1=Borror |first1=D. J. |first2=Dwight M. |last2=DeLong |first3=Charles A. |last3=Triplehorn |title=Introduction to the Study of Insects |location=New York |publisher=Holt, Rinehart & Winston |edition=Sixth |year=2004 |isbn=0-03-096835-6 }}</ref>
+The pupae of different groups of insects have different names such as [[#Chrysalis|''chrysalis'']] for the pupae of [[butterflies]] and ''tumbler'' for those of the [[mosquito]] family. Pupae may further be enclosed in other structures such as [[#Cocoon|cocoons]], [[nest]]s, or [[Exoskeleton|shells]].<ref>{{cite book |last1=Borror |first1=D. J. |first2=Dwight M. |last2=DeLong |first3=Charles A. |last3=Triplehorn |title=Introduction to the Study of Insects |location=New York |publisher=Holt, Rinehart & Winston |edition=Sixth |year=2004 |isbn=0-03-096835-6 }}</ref>
 ==Position in life cycle==
-The pupal stage follows the [[larva]]l stage and precedes [[adult]]hood (''[[imago]]'') in insects with complete metamorphosis. The pupa is a non-feeding, usually [[Sessility (zoology)|sessile]] stage, or highly active as in mosquitoes. It is during the pupal stage that the adult structures of the insect are formed while the larval structures are broken down. The adult structures grow from [[imaginal disc]]s.<ref name="AldazEscudero2010">{{cite journal |last1=Aldaz |first1=Silvia |last2=Escudero |first2=Luis M. |title=Imaginal discs |journal=Current Biology |date=2010 |volume=20 |issue=10 |pages=R429–R431 |doi=10.1016/j.cub.2010.03.010|pmid=20504747 |doi-access=free }}</ref>
+The pupal stage follows the [[larva]]l stage, or in some cases a [[Prepupa|prepupal]] stage, and precedes [[adult]]hood (''[[imago]]'') in insects with complete metamorphosis. The pupa is a non-feeding, usually [[Sessility (zoology)|sessile]] stage, or highly active as in mosquitoes. It is during the pupal stage that the adult structures of the insect are formed while the larval structures are broken down. The adult structures grow from [[imaginal disc]]s.<ref name="AldazEscudero2010">{{cite journal |last1=Aldaz |first1=Silvia |last2=Escudero |first2=Luis M. |title=Imaginal discs |journal=Current Biology |date=2010 |volume=20 |issue=10 |pages=R429–R431 |doi=10.1016/j.cub.2010.03.010|pmid=20504747 |doi-access=free |bibcode=2010CBio...20.R429A }}</ref>
 ===Duration===
@@ Line 17: / Line 17: @@
 ===Emergence===
-Insects emerge (eclose) from pupae by splitting the pupal case. Most butterflies emerge in the morning. In mosquitoes, the emergence is in the evening or night. In fleas, the process is triggered by vibrations that indicate the possible presence of a suitable host. Prior to emergence, the adult inside the pupal [[exoskeleton]] is termed ''pharate''. Once the pharate adult has eclosed from the pupa, the empty pupal exoskeleton is called an ''[[exuvia]]''; in most [[hymenopteran]]s (ants, bees and wasps) the exuvia is so thin and membranous that it becomes "crumpled" as it is shed.
+Insects emerge (eclose) from pupae by splitting the pupal case. Most butterflies emerge in the morning. In mosquitoes, the emergence is in the evening or night. In fleas, the process is triggered by vibrations that indicate the possible presence of a suitable host. Prior to emergence, the adult inside the pupal [[exoskeleton]] is termed ''pharate''. Once the pharate adult has eclosed from the pupa, the empty pupal exoskeleton is called an ''[[exuvia]]''; in most [[hymenopteran]]s (ants, bees and wasps) the exuvia is so thin and membranous that it becomes "crumpled" as it is shed. Measuring the timing of this emergence is of interest to chronobiologists because the process is regulated by [[Circadian clock|circadian clocks]] in many species, necessitating different [[Eclosion assay|assays]] to measure eclosion timing.
 ===Pupal mating===
@@ Line 24: / Line 24: @@
 ==Defense==
-Pupae are usually immobile and are largely defenseless. To overcome this, pupae often are covered with a cocoon, conceal themselves in the environment, or form underground.<ref>{{Cite book|title=The insects : structure and function|last=Chapman, R. F. (Reginald Frederick)|year=2013|isbn=978-0-521-11389-2|pages=419|oclc=794624696}}</ref> There are some species of [[Lycaenidae|Lycaenid butterflies]] which are protected in their pupal stage by ants. Another means of defense by pupae of other species is the capability of making sounds or vibrations to scare potential predators. A few species use chemical defenses including toxic secretions. The pupae of social [[hymenoptera]]ns are protected by adult members of the hive.
+Pupae are usually immobile and are largely defenseless. To overcome this, pupae often are covered with a cocoon, conceal themselves in the environment, or form underground.<ref>{{Cite book|title=The insects : structure and function|last=Chapman, R. F. (Reginald Frederick)|year=2013|isbn=978-0-521-11389-2|pages=419|publisher=Cambridge University Press |oclc=794624696}}</ref> Some species of [[Lycaenidae|Lycaenid butterflies]] are protected in their pupal stage by ants. Another means of defense by pupae of other species is the capability of making sounds or vibrations to scare potential predators. A few species use chemical defenses including toxic secretions. The pupae of social [[hymenoptera]]ns are protected by adult members of the hive.
 ==Types==
-Based on the presence or absence of articulated [[mandible]]s that are employed in emerging from a cocoon or pupal case, the pupae can be classified in to two types:<ref>{{cite web | url=http://www.agriinfo.in/default.aspx?page=topic&superid=6&topicid=1553 | title=Types of Pupa | publisher=Agri Info | access-date=28 April 2016 | archive-url=https://web.archive.org/web/20170204080242/http://www.agriinfo.in/default.aspx?page=topic&superid=6&topicid=1553 | archive-date=4 February 2017 | url-status=dead }}</ref><ref>{{cite web | url=http://what-when-how.com/insects/pupa-and-puparium-insects/ | title=Pupa and Puparium (Insects) | publisher=what-when-how | access-date=28 April 2016 | archive-url=https://web.archive.org/web/20180122141431/http://what-when-how.com/insects/pupa-and-puparium-insects/ | archive-date=22 January 2018 | url-status=dead }}</ref>
+Based on the presence or absence of articulated [[mandible]]s that are employed in emerging from a cocoon or pupal case, the pupae can be classified in to two types:<ref>{{cite web | url=http://www.agriinfo.in/default.aspx?page=topic&superid=6&topicid=1553 | title=Types of Pupa | publisher=Agri Info | access-date=28 April 2016 | archive-url=https://web.archive.org/web/20170204080242/http://www.agriinfo.in/default.aspx?page=topic&superid=6&topicid=1553 | archive-date=4 February 2017 | url-status=dead }}</ref>
 * '''Decticous pupa''' – a pupa with articulated mandibles. Examples are pupae of the orders [[Neuroptera]], [[Mecoptera]], [[Trichoptera]] and few [[Lepidoptera]] families.
 * '''Adecticous pupa''' – a pupa without articulated mandibles. Examples include the orders [[Strepsiptera]], [[Coleoptera]], [[Hymenoptera]], [[Diptera]] and [[Siphonaptera]].
-Based on whether the pupal appendages are free or attached to the body, the pupae can be classified as one of three types:<ref>{{cite web | url=http://insects.about.com/od/morphology/tp/Insect-Pupal-Forms.htm | title=Insect Pupal Forms | publisher=About.com | access-date=28 April 2016}}</ref>
+Based on whether the pupal appendages are free or attached to the body, the pupae can be classified as one of three types:<!--<ref>{{cite web | url=http://insects.about.com/od/morphology/tp/Insect-Pupal-Forms.htm | title=Insect Pupal Forms | publisher=About.com | access-date=28 April 2016}}</ref>-->
 * '''Exarate pupa''' – appendages are free and are not usually encapsulated within a cocoon. Decticous pupae are always exarate; some adecticous pupae are as well. (Neuroptera, Trichoptera, [[Cyclorrhapha]] of Dipterans, Siphonaptera, most Coleoptera, Hymenoptera, and few Lepidoptera).
 * '''Obtect pupa''' – appendages are attached closely to the body and are commonly encapsulated within a cocoon. Some adecticous pupa are obtect forms. (Most Lepidoptera, [[Nematocera]] and [[Brachycera]] of Dipterans, [[Staphylinidae]] and [[Chrysomelidae]] Coleopterans, many [[Chalcidoidea]] Hymenopterans)
@@ Line 38: / Line 38: @@
 ==Chrysalis==
 {{Redirect|Chrysalis}}
-[[Image:Common crow pupa.jpg|thumb|200px|Common crow butterfly (''[[Euploea core]]'') chrysalis illustrating the [[Ancient Greek language|Ancient Greek]] origin of the term: {{lang|grc|χρυσός}} ({{transl|grc|chrysós}}) for gold]]
+[[File:A pupa of Euploea core (Cramer, 1780) - Common Crow WLB.jpg|thumb|Common crow butterfly (''[[Euploea core]]'') chrysalis illustrating the [[Ancient Greek language|Ancient Greek]] origin of the term: {{lang|grc|χρυσός}} ({{transl|grc|chrysós}}) for gold]]
-[[File:Painted Lady Chrysalis micro CT.jpg|thumb|200px|[[X-ray microtomography|Micro CT]] image showing inside chrysalis of a painted lady butterly (''[[Vanessa cardui]]'')]]
 A chrysalis ({{lang-la|chrysallis}}, from {{lang-grc|χρυσαλλίς}}, {{transl|grc|chrysallís}}, plural: {{transl|grc|chrysalides}}, also known as an ''aurelia'') or nympha is the pupal stage of [[butterflies]]. The term is derived from the metallic–gold coloration found in the pupae of many butterflies, referred to by the Ancient Greek term {{lang|grc|χρυσός}} ({{transl|grc|chrysós}}) for gold.
@@ Line 45: / Line 44: @@
 When the caterpillar is fully grown, it makes a button of silk which it uses to fasten its body to a leaf or a twig. Then the caterpillar's skin comes off for the final time. Under this old skin is a hard skin called a chrysalis.<ref>{{cite book |last=Darby |first=Gene |title=What is a Butterfly |url=https://archive.org/details/whatisbutterfly00darb |url-access=registration |publisher=[[Benefic Press]] |location=Chicago |year=1958 |page=[https://archive.org/details/whatisbutterfly00darb/page/19 19]}}</ref>
-Because chrysalises are often showy and are formed in the open, they are the most familiar examples of pupae. Most chrysalides are attached to a surface by a [[Hook and loop fastener|Velcro]]-like arrangement of a [[silk]]en pad spun by the [[caterpillar]], usually cemented to the underside of a perch, and the ''cremastral'' hook or hooks protruding from the rear of the chrysalis or ''cremaster'' at the tip of the pupal [[abdomen]] by which the caterpillar fixes itself to the pad of silk. ({{abbr|Gr.|Ancient Greek}} {{transl|grc|kremastos}} 'suspended')<ref>[http://en.academic.ru/dic.nsf/en_invertebrate_zoology/3481/cremaster Academic Dictionaries and Encyclopedias]</ref>
+Because chrysalises are often showy and are formed in the open, they are the most familiar examples of pupae. Most chrysalides are attached to a surface by a [[Hook and loop fastener|Velcro]]-like arrangement of a [[silk]]en pad spun by the [[caterpillar]], usually cemented to the underside of a perch, and the ''cremastral'' hook or hooks protruding from the rear of the chrysalis or ''cremaster'' at the tip of the pupal [[abdomen]] by which the caterpillar fixes itself to the pad of silk. ({{abbr|Gr.|Ancient Greek}} {{transl|grc|kremastos}} 'suspended')<ref>{{Cite web |url=http://en.academic.ru/dic.nsf/en_invertebrate_zoology/3481/cremaster |title=Academic Dictionaries and Encyclopedias |access-date=2013-01-20 |archive-date=2013-05-25 |archive-url=https://web.archive.org/web/20130525045759/http://en.academic.ru/dic.nsf/en_invertebrate_zoology/3481/cremaster |url-status=live }}</ref>
 Like other types of pupae, the chrysalis stage in most butterflies is one in which there is little movement. However, some butterfly pupae are capable of moving the abdominal segments to produce sounds or to scare away potential [[predator]]s. Within the chrysalis, growth and differentiation occur.<ref name=Lowe>{{cite journal|title=Metamorphosis revealed: three dimensional imaging inside a living chrysalis|first1=Tristan|last1=Lowe|first2=Russell P.|last2=Garwood|first3=Thomas|last3=Simonsen|first4=Robert S.|last4=Bradley|first5=Philip J.|last5=Withers|year=2013|journal=Journal of the Royal Society Interface |volume=10|issue=84|at=20130304|doi=10.1098/rsif.2013.0304|pmid=23676900|pmc=3673169}}</ref> The adult butterfly emerges (ecloses) from this and expands its wings by pumping [[haemolymph]] into the wing veins.<ref>[http://www.amnh.org/exhibitions/butterflies/highlights/metamorphosis.php AMNH] {{webarchive|url=https://web.archive.org/web/20061207003856/http://amnh.org/exhibitions/butterflies/highlights/metamorphosis.php |date=December 7, 2006}} Accessed December 2006</ref> Although this sudden and rapid change from pupa to imago is often called [[metamorphosis]], metamorphosis is really the whole series of changes that an insect undergoes from egg to adult.
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 When emerging, the butterfly uses a liquid, sometimes called [[cocoonase]], which softens the shell of the chrysalis. Additionally, it uses two sharp claws located on the thick joints at the base of the forewings to help make its way out.<ref>[https://archive.org/stream/entomologist351902brit/entomologist351902brit_djvu.txt ''The Entomologist'']</ref> Having emerged from the chrysalis, the butterfly will usually sit on the empty shell in order to expand and harden its wings. However, if the chrysalis was near the ground (such as if it fell off from its silk pad), the butterfly would find another vertical surface to rest upon and harden its wings (such as a wall or fence).
-[[Moth]] pupae are usually dark in color and either formed in underground cells, loose in the soil, or their pupa is contained in a protective [[silk]] case called a ''cocoon''. The pupa of some species such as the [[hornet moth]] develop sharp ridges around the outside called adminicula that allow the pupa to move from its place of concealment inside a tree trunk when it is time for the adult to emerge.<ref>{{Cite journal|last=Newland|first=D.E.|title=Eclosion mechanics, mating and ovipositing behaviour of ''Sesia apiformis''|journal=Entomologist's Gazette}}</ref>
+[[Moth]] pupae are usually dark in color and either formed in underground cells, loose in the soil, or their pupa is contained in a protective [[silk]] case called a ''cocoon''. The pupa of some species such as the [[hornet moth]] develops sharp ridges around the outside called adminicula that allow the pupa to move from its place of concealment inside a tree trunk when it is time for the adult to emerge.<ref>{{Cite journal|last=Newland|first=D.E.|title=Eclosion mechanics, mating and ovipositing behaviour of ''Sesia apiformis''|journal=Entomologist's Gazette}}</ref>
 Pupa, chrysalis, and cocoon are frequently confused, but are quite distinct from each other. The pupa is the stage between the larva and adult stages. The chrysalis generally refers to a butterfly pupa although the term may be misleading as there are some moths whose pupae resembles a chrysalis, e.g.: the plume winged moths of the family [[Pterophoridae]] and some [[geometrid]] moths. A cocoon is a silk case that the larvae of moths, and sometimes other insects, spin around the pupa.
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 ==Cocoon==
 {{Other uses|Cocoon (disambiguation)}}
+[[File:Dwarf birch spinner (Eriogaster arbusculae) cocoon Dovrefjell.jpg|thumb|left|Dwarf birch spinner moth (''[[Eriogaster arbusculae]]'') cocoon]]
-[[Image:Caterpillars cocoon.jpg|thumb|The tough brown cocoon of an [[Opodiphthera eucalypti|emperor gum moth]]]]
 A cocoon is a casing spun of [[silk]] by many [[moth]]s and [[caterpillar]]s,<ref>{{Cite book | last = Darby | first = Gene | title = What is a Butterfly | publisher = Benefic Press | location = Chicago | year = 1958 | page = 41 }}</ref> and numerous other [[holometabolous]] insect [[larva]]e as a protective covering for the pupa.
-Cocoons may be tough or soft, opaque or translucent, solid or meshlike, of various colors, or composed of multiple layers, depending on the type of insect larva producing it. Many moth caterpillars shed the larval hairs ([[seta]]e) and incorporate them into the cocoon; if these are [[urticating hair]]s then the cocoon is also irritating to the touch. Some larvae attach small twigs, fecal pellets or pieces of vegetation to the outside of their cocoon in an attempt to disguise it from predators. Others spin their cocoon in a concealed location—on the underside of a [[leaf]], in a [[Fracture (geology)|crevice]], down near the base of a [[tree trunk]], suspended from a twig or concealed in the [[leaf litter]].<ref>{{cite book |first=Malcolm J. |last=Scoble |year=1992 |title=The Lepidoptera: Form, Function and Diversity |url=https://archive.org/details/lepidopteraformf0000scob |url-access=registration |location=Oxford |publisher=Oxford University Press |isbn=0-19-854031-0 }}</ref>
+Cocoons may be tough or soft, opaque or translucent, solid or meshlike, of various colors, or composed of multiple layers, depending on the type of insect larva producing it. Many moth caterpillars shed the larval hairs ([[seta]]e) and incorporate them into the cocoon; if these are [[urticating hair]]s then the cocoon is also irritating to the touch. Some larvae attach small twigs, fecal pellets or pieces of vegetation to the outside of their cocoon in an attempt to disguise it from predators. Others spin their cocoon in a concealed location—on the underside of a [[leaf]], in a [[Fracture (geology)|crevice]], down near the base of a [[tree trunk]], suspended from a twig or concealed in the [[leaf litter]].<ref>{{cite book |first=Malcolm J. |last=Scoble |year=1992 |title=The Lepidoptera: Form, Function and Diversity |url=https://archive.org/details/lepidopteraformf0000scob |url-access=registration |location=Oxford |publisher=Oxford University Press |isbn=0-19-854031-0 }}</ref> Contrary to popular belief, larvae do not completely liquify inside the cocoon.<ref>{{Cite news |last=Guarino |first=Ben |date=September 19, 2017 |title=Mutant butterflies reveal the genetic roots of colorful wings |url=https://www.washingtonpost.com/news/speaking-of-science/wp/2017/09/19/mutant-butterflies-reveal-the-genetic-roots-of-colorful-wings/ |url-status=live |archive-url=https://web.archive.org/web/20170919133746/https://www.washingtonpost.com/web/20170919133746/https://www.washingtonpost.com/news/speaking-of-science/wp/2017/09/19/mutant-butterflies-reveal-the-genetic-roots-of-colorful-wings/?utm_term=.6a605ce29ece |archive-date=September 19, 2017 |access-date=July 6, 2024 |work=[[The Washington Post]]}}</ref>
+[[Image:Caterpillars cocoon.jpg|thumb|upright|The tough brown cocoon of an [[Opodiphthera eucalypti|emperor gum moth]]]]
-The silk in the cocoon of the [[silk moth]] can be unraveled to harvest silk fibre which makes this moth the most economically important of all lepidopterans. The silk moth is the only completely domesticated lepidopteran and does not exist in the wild.
+The silk in the cocoon of the [[Bombyx mori|silk moth]] can be unraveled to harvest silk fibre which makes this moth the most economically important of all lepidopterans. The silk moth is the only completely domesticated lepidopteran; it does not exist in the wild.
 Insects that pupate in a cocoon must escape from it, and they do this either by the pupa cutting its way out, or by secreting [[enzyme]]s, sometimes called [[cocoonase]], that soften the cocoon. Some cocoons are constructed with built-in lines of weakness along which they will tear easily from inside, or with exit holes that only allow a one-way passage out; such features facilitate the escape of the adult insect after it emerges from the pupal skin.
+==Puparium==
 [[File:Eupeodes americanus.jpg|thumb|Puparium of ''[[Eupeodes americanus]]'']]
+Some pupae remain inside the exoskeleton of the final larval [[instar]] and this last larval "shell" is called a puparium (plural, puparia). Flies of the group [[Muscomorpha]] have puparia, as do members of the order [[Strepsiptera]], and the [[Hemiptera]]n family [[Aleyrodidae]].<ref>{{cite book |last1=Gullan |first1=P. J. |last2=Cranston |first2=P. S. |year=2000 |title=The Insects: An Outline of Entomology |edition=2nd |location=London |publisher=Blackwell |isbn=0-632-05343-7 }}</ref>
-==Puparium==
-Some pupae remain inside the exoskeleton of the final larval [[instar]] and this last larval "shell" is not called a puparium (plural, puparia). Flies of the group [[Muscomorpha]] have puparia, as do members of the order [[Strepsiptera]], and the [[Hemiptera]]n family [[Aleyrodidae]].<ref>{{cite book |last1=Gullan |first1=P. J. |last2=Cranston |first2=P. S. |year=2000 |title=The Insects: An Outline of Entomology |edition=2nd |location=London |publisher=Blackwell |isbn=0-632-05343-7 }}</ref>
 ==Gallery==
-<gallery mode="packed" heights="100">
+<gallery >
+File:Rhynchophorus ferrugineus (pupa)-2.jpg|''[[Rhynchophorus ferrugineus]]'' pupa
 File:Fruit Fly Pupa.jpg|Fruit fly (''[[Drosophila melanogaster]]'') pupa
 File:Caterpillar making cocoon2.jpg|An [[emperor gum moth]] caterpillar spinning its cocoon
 File:Actias luna pupa 2 sjh.JPG|[[Actias luna|Luna moth]] cocoon and pupa
-File:Actias luna cocoons sjh.JPG|Assortment of Luna moth cocoons
+File:Actias luna cocoons sjh.JPG|Assortment of luna moth cocoons
 File:Actias luna emergence sjh.gif|Luna moth emerging from pupa within [[silk cocoon]]
 File:Actias luna pupa sjh enhanced.JPG|Luna moth pupa removed from cocoon
@@ Line 83: / Line 83: @@
 File:Pupation - Inachis io.jpg|Pupation of ''[[Aglais io]]''
 File:Monarch Butterfly Chrysalis.JPG|[[Monarch butterfly]] chrysalis
-File:Nacimiento de una Dryas iulia, Mariposario de Icod de los Vinos, Tenerife, España, 2012-12-13, DD 03.jpg|Specimen of an [[Eclosion|eclosing]] ''[[Dryas iulia]]'' butterfly
+File:Nacimiento de una Dryas iulia, Mariposario de Icod de los Vinos, Tenerife, España, 2012-12-13, DD 03.jpg|Specimen of an eclosing ''[[Dryas iulia]]'' butterfly
-File:Pupae of Japanagromyza inferna Spencer in gall of Centrosema virginianum L. - ZooKeys-374-045-g005.jpg|Pupae of ''Japanagromyza inferna'', a [[gall fly]], in [[gall]] of ''[[Centrosema virginianum]]''
+File:Pupae of Japanagromyza inferna Spencer in gall of Centrosema virginianum L. - ZooKeys-374-045-g005.jpg|Pupae of ''[[Japanagromyza inferna]]'', a [[gall fly]], in [[gall]] of ''[[Centrosema virginianum]]''
 File:Cabbage looper in cocoon.jpg|Pupa of [[cabbage looper]]
-File:BARON BUTTERFLY CHRYSALIS.jpg|Pupa of Baron Butterfly [[Euthalia aconthea]]
+File:BARON BUTTERFLY CHRYSALIS.jpg|Pupa of baron butterfly ''[[Euthalia aconthea]]''
-File:Papilio polytes common Mormon pupa vijayanrajapuram 03.jpg|thumb|Papilio_polytes_common_Mormon_pupa on curry leaves.
+File:Papilio polytes common Mormon pupa vijayanrajapuram 03.jpg|''[[Papilio polytes]]'' common Mormon pupa on curry leaves.
+File:Annual cicada pupating 2024-08-12 No 2.jpg|''[[Dog-day cicada]]'' emerging
 </gallery>
 ==See also==
-* [[Brood (honeybee)]]
+* [[Bee brood]]
 * [[Larva]]
 * [[Silk]]
@@ Line 101: / Line 102: @@
 {{Commons category|Pupae}}
 * [http://insecta.pro/gallery?fem=5 Pupa photos] plus species descriptions at ''Insecta.pro''
-* [https://archive.today/20121108172719/http://picasaweb.google.com/srither240255/CycleOfSilkworm Silk worm life cycle photos]
+* [https://archive.today/20121108172719/http://picasaweb.google.com/srither240255/CycleOfSilkworm Silk worm life cycle photos] (archived 8 November 2012)
 {{Authority control}}